Clathrus ruber
Encyclopedia
Clathrus ruber is a species of fungus
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...

 in the stinkhorn family, and the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 of the genus Clathrus
Clathrus
Clathrus is a genus of basidiomycete fungi of the family Phallaceae. As with other stinkhorn fungi, mature fruit bodies are covered with olive-brown slimy gleba, containing spores, which attracts flies...

. It is commonly
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 known as the latticed stinkhorn, the basket stinkhorn, or the red cage, alluding to the striking fruit bodies
Sporocarp (fungi)
In fungi, the sporocarp is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne...

 that are shaped somewhat like a round or oval hollow sphere with interlaced or latticed branches. The fungus is saprobic, feeding off decaying woody plant material, and is usually found alone or in groups in leaf litter on garden soil, grassy places, or on woodchip
Woodchipping
Woodchipping is the act and industry of chipping wood for pulp, processed wood products, and mulch.-Papermaking:Timber is converted to woodchips and sold, primarily, for pulp production used in paper manufacture...

 garden mulch
Mulch
In agriculture and gardening, is a protective cover placed over the soil to retain moisture, reduce erosion, provide nutrients, and suppress weed growth and seed germination. Mulching in gardens and landscaping mimics the leaf cover that is found on forest floors....

es. Although considered primarily a European species, C. ruber has been introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 to other areas, and now has a wide distribution that includes northern Africa, Asia, Australia, and North and South America. The species was illustrated in the scientific literature during the 1500s, but was not officially described
Species description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...

 until 1729.

The fruit body
Basidiocarp
In fungi, a basidiocarp, basidiome or basidioma , is the sporocarp of a basidiomycete, the multicellular structure on which the spore-producing hymenium is borne. Basidiocarps are characteristic of the hymenomycetes; rusts and smuts do not produce such structures...

 initially appears like a whitish "egg" attached to the ground at the base by cords called rhizomorphs. The egg has a delicate, leathery outer membrane enclosing the compressed lattice that surrounds a layer of olive-green spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

-bearing slime called the gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

, which contains high levels of calcium
Calcium
Calcium is the chemical element with the symbol Ca and atomic number 20. It has an atomic mass of 40.078 amu. Calcium is a soft gray alkaline earth metal, and is the fifth-most-abundant element by mass in the Earth's crust...

 that help protect the developing fruit body during development. As the egg ruptures and the fruit body expands, the gleba is carried upward on the inner surfaces of the spongy lattice, and the egg membrane remains as a volva
Volva (mycology)
The volva is a mycological term to describe a cup-like structure at the base of a mushroom that is a remnant of the universal veil. This macrofeature is important in wild mushroom identification due to it being an easily observed, taxonomically significant feature which frequently signifies a...

 around the base of the structure. The fruit body can reach heights of up to 20 cm (7.9 in). The color of the fruit body, which can range from pink to orange to red, results primarily from the carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

 pigments lycopene
Lycopene
Lycopene is a bright red carotene and carotenoid pigment and phytochemical found in tomatoes and other red fruits and vegetables, such as red carrots, watermelons and papayas...

 and beta-carotene. The gleba has a fetid odor, somewhat like rotting meat
Carrion
Carrion refers to the carcass of a dead animal. Carrion is an important food source for large carnivores and omnivores in most ecosystems. Examples of carrion-eaters include vultures, hawks, eagles, hyenas, Virginia Opossum, Tasmanian Devils, coyotes, Komodo dragons, and burying beetles...

, which attracts flies and other insects to help disperse its spores. Although the edibility
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

 of the fungus is not known with certainty, its odor would deter most from consuming it. C. ruber was not regarded highly in tales in southern European folklore, which suggested that those who handled the mushroom risked contracting various ailments.

Taxonomy, phylogeny, and naming

Clathrus ruber was illustrated as early as 1560 by the Swiss naturalist Conrad Gesner in his Nomenclator Aquatilium Animantium—Gesner mistook the mushroom for a marine organism. It appeared in a woodcut
Woodcut
Woodcut—occasionally known as xylography—is a relief printing artistic technique in printmaking in which an image is carved into the surface of a block of wood, with the printing parts remaining level with the surface while the non-printing parts are removed, typically with gouges...

 in John Gerard
John Gerard
John Gerard aka John Gerarde was an English herbalist notable for his herbal garden and botany writing. In 1597 he published a large and heavily illustrated "Herball, or Generall Historie of Plantes", which went on to be the most widely circulated botany book in English in the 17th century...

's 1597 Great Herball, shortly thereafter in Carolus Clusius 1601 Fungorum in Pannoniis Observatorum Brevis Historia, and was one of the species featured in Cassiano dal Pozzo
Cassiano dal Pozzo
Cassiano dal Pozzo was an Italian scholar and patron of arts. The secretary of Cardinal Francesco Barberini, he was an antiquary in the classicizing circle of Rome, and a long-term friend and patron of Nicolas Poussin, whom he supported from his earliest arrival in Rome: Poussin in a letter...

's museo cartaceo ("paper museum") that consisted of thousands of illustrations of the natural world.

The fungus was first described
Species description
A species description or type description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species which have been described previously, or are...

 scientifically in 1729, by the Italian Pier Antonio Micheli
Pier Antonio Micheli
Pier Antonio Micheli was a noted Italian botanist, professor of botany in Pisa, curator of the Orto Botanico di Firenze, author of Nova plantarum genera iuxta Tournefortii methodum disposita...

 in his Nova plantarum genera iuxta Tournefortii methodum disposita, who gave it its current scientific name. The species was once referred to by American authors as Clathrus cancellatus L., as they used a system of nomenclature
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....

 based on the former American Code of Botanical Nomenclature, in which the starting point for naming species was Linnaeus's 1753 Species Plantarum
Species Plantarum
Species Plantarum was first published in 1753, as a two-volume work by Carl Linnaeus. Its prime importance is perhaps that it is the primary starting point of plant nomenclature as it exists today. This means that the first names to be considered validly published in botany are those that appear...

. The International Code for Botanical Nomenclature now uses the same starting date, but names of Gasteromycetes used by Christian Hendrik Persoon
Christian Hendrik Persoon
Christiaan Hendrik Persoon was a mycologist who made additions to Linnaeus' mushroom taxonomy.-Early life:...

 in his Synopsis Methodica Fungorum (1801) are sanctioned
Sanctioned name
In mycology, a sanctioned name is a name that was adopted in certain works of Christiaan Hendrik Persoon or Elias Magnus Fries, which are considered major points in fungal taxonomy.-Definition and effects:...

 and automatically replace earlier names. Since Persoon used the specific epithet ruber, the correct name for the species is Clathrus ruber. Several historical names of the fungus are now synonyms
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...

: Clathrus flavescens, named by Persoon in 1801; Clathrus cancellatus by Joseph Pitton de Tournefort
Joseph Pitton de Tournefort
Joseph Pitton de Tournefort was a French botanist, notable as the first to make a clear definition of the concept of genus for plants.- Biography :...

 and published by Elias Fries in 1823; Clathrus nicaeensis, published by Jean-Baptiste Barla
Jean-Baptiste Barla
Jean-Baptiste Barla was a French botanist.Barla was a man of independent means and dedicated himself to investigate the fungi and orchids of the South of France. "Cette végétation exceptionelle, doit évidemment sa richesse et sa variété à la configuration topographique, toute particulière, du...

 in 1879; and Clathrus ruber var. flavescens, published by Livio Quadraccia and Dario Lunghini in 1990.

Clathrus ruber is the type species
Type species
In biological nomenclature, a type species is both a concept and a practical system which is used in the classification and nomenclature of animals and plants. The value of a "type species" lies in the fact that it makes clear what is meant by a particular genus name. A type species is the species...

 of the genus Clathrus
Clathrus
Clathrus is a genus of basidiomycete fungi of the family Phallaceae. As with other stinkhorn fungi, mature fruit bodies are covered with olive-brown slimy gleba, containing spores, which attracts flies...

, and is part of the group of Clathrus species known as the Laternoid series. Common features uniting this group include the vertical arms of the receptacle (fruit body) that are not joined together at the base, and the spongy structure of the receptacle. According to a molecular analysis published in 2006, out of the about 40 Phallales species used in the study, C. ruber is most closely related to Aseroe rubra
Aseroë rubra
Aseroe rubra, commonly known as the anemone stinkhorn, sea anemone fungus and starfish fungus, is a common and widespread basidiomycete fungus recognizable for its foul odour of carrion and its sea anemone shape when mature. Found in gardens on mulch and in grassy areas, it resembles a red...

, Clathrus archeri
Clathrus archeri
Clathrus archeri , commonly known as Octopus Stinkhorn, is indigenous to Australia and Tasmania and an introduced species in Europe and North America. The young fungus erupts from a suberumpent egg by forming into four to seven elongated slender arms initially erect and attached at the top...

, Laternea triscapa
Laternea triscapa
Laternea triscapa is a species of fungi in the Phallaceae family. The species was first described by French botanist Pierre Jean François Turpin in 1820. It is found in Central and South America, and the West Indies....

, and Clathrus chrysomycelinus
Clathrus chrysomycelinus
Clathrus chrysomycelinus is a species of fungus in the stinkhorn family. It is found in South America....

.

The generic name Clathrus is derived from Ancient Greek κλειθρον or "lattice", and the specific epithet is Latin ruber, meaning "red". The mushroom is commonly
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 known as the "basket stinkhorn", the "lattice stinkhorn", or the "red cage". It was known to the locals of the Adriatic hinterland in the former Yugoslavia
Former Yugoslavia
The former Yugoslavia is a term used to describe the present day states which succeeded the collapse of the Socialist Federal Republic of Yugoslavia....

 as veštičije srce or vještičino srce, meaning "witch's heart".

Description

Before the volva
Volva (mycology)
The volva is a mycological term to describe a cup-like structure at the base of a mushroom that is a remnant of the universal veil. This macrofeature is important in wild mushroom identification due to it being an easily observed, taxonomically significant feature which frequently signifies a...

 opens, the fruiting body is egg-shaped to roughly spherical, up to 6 cm (2.4 in) in diameter, with a gelatinous interior up to 3 mm (0.118110236220472 in) thick. White to grayish in color, it is initially smooth, but develops a network of polygonal marks on the surface prior to opening as the internal structures expand and stretch the peridium
Peridium
The peridium is the protective layer that encloses a mass of spores in fungi. This outer covering is a distinctive feature of the Gasteromycetes.-Description:...

 taut. The fruit body, or receptacle, bursts the egg open as it expands (a process that can take as little as a few hours), and leaves the remains of the peridium as a cup or volva
Volva (mycology)
The volva is a mycological term to describe a cup-like structure at the base of a mushroom that is a remnant of the universal veil. This macrofeature is important in wild mushroom identification due to it being an easily observed, taxonomically significant feature which frequently signifies a...

 surrounding the base. The receptacle ranges in color from red to bright pink to pale orange, and it is often lighter in color approaching the base. The color appears to be dependent upon the temperature and humidity of the environment. The receptacle consists of a spongy network of "arms" interlaced to make meshes of unequal size. At the top of the receptacle, the arms are up to 1.5 cm (0.590551181102362 in) thick, but they taper down to smaller widths near the base. A cross-section of the arm reveals it to be spongy, and made up of one wide inner tube and two indistinct rows of tubes towards the outside. The outer surface of the receptacle is ribbed or wrinkled. There are between 80 and 120 mesh holes in the receptacle. The unusual shape of the receptacle has inspired some creative comparisons: David Arora
David Arora
David Arora is an American mycologist, naturalist, and writer. He is the author of two popular books on mushroom identification, Mushrooms Demystified and All That the Rain Promises and More.......

 likened it to a whiffleball, while the German Mycological Society—who named C. ruber the 2011 "Mushroom of the Year"—described it as "like an alien from a science fiction horror film".

A considerable variation in height has been reported for the receptacle, ranging from 8 to 20 cm (3.1 to 7.9 in) tall. The base of the fruit bodies are attached to the substrate
Substrate (biology)
In biology a substrate is the surface a plant or animal lives upon and grows on. A substrate can include biotic or abiotic materials and animals. For example, encrusting algae that lives on a rock can be substrate for another animal that lives on top of the algae. See also substrate .-External...

 by rhizomorphs (thickened cords of mycelia). The dark olive-green to olive-brown, foul-smelling sticky gleba
Gleba
Gleba is the fleshy spore-bearing inner mass of fungi such as the puffball or stinkhorn.The gleba is a solid mass of spores, generated within an enclosed area within the sporocarp. The continuous maturity of the sporogenous cells leave the spores behind as a powdery mass that can be easily blown away...

 covers the inner surface of the receptacle, except near the base. The odor—described as resembling rotting meat—attracts flies, other insects, and, in one report, a scarab beetle (Scarabaeus sacer
Scarabaeus sacer
Scarabaeus sacer is a species of dung beetle, found in coastal dunes and marshes around the Mediterranean Basin. It collects balls of dung which it rolls to an underground chamber to feed its offspring. This behaviour inspired the Ancient Egyptians to compare it to the sun god Khepri, and they...

) that help disperse the spores. The putrid odor—and people's reaction to it—have been well documented. In 1862 Mordecai Cubitt Cooke
Mordecai Cubitt Cooke
Mordecai Cubitt Cooke was an English botanist and mycologist.Cooke came from a mercantile family in Horning, Norfolk, and worked as an apprentice to a fabric merchant before becoming a clerk in a law firm, but his chief interest was in botany. He founded the Society of Amateur Botanists in 1862...

 wrote "it is recorded of a botanist who gathered one for the purpose of drying it for his herbarium, that he was compelled by the stench to rise during the night and cast the offender out the window." American mycologist David Arora
David Arora
David Arora is an American mycologist, naturalist, and writer. He is the author of two popular books on mushroom identification, Mushrooms Demystified and All That the Rain Promises and More.......

 called the odor "the vilest of any stinkhorn". The receptacle collapses about 24 hours after its initial eruption from the egg.

The spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s are elongated, smooth, and have dimensions of 4–6 by 1.5–2 µm
Micrometre
A micrometer , is by definition 1×10-6 of a meter .In plain English, it means one-millionth of a meter . Its unit symbol in the International System of Units is μm...

. Scanning electron microscopy has revealed that C. ruber (in addition to several other Phallales species) has a hilar
Hilum (biology)
In botany, the hilum is a scar or mark left on a seed coat by the former attachment to the ovary wall or to the funiculus...

 scar—a small indentation in the surface of the spore where it was previously connected to the basidium
Basidium
thumb|right|500px|Schematic showing a basidiomycete mushroom, gill structure, and spore-bearing basidia on the gill margins.A basidium is a microscopic, spore-producing structure found on the hymenophore of fruiting bodies of basidiomycete fungi. The presence of basidia is one of the main...

 via the sterigma
Sterigma
thumb|right|The sterigmata is the slender extension that connects the spore to the basidia .A sterigma is an extension of the basidium consisting of a basal filamentous part and a slender projection with a spore at the tip...

. The basidia (spore-bearing cells) are six-spored.

Similar species

Clathrus ruber may be distinguished from the closely related tropical species C. crispus
Clathrus crispus
Clathrus crispus is a species of fungus in the stinkhorn family. It is found in the Americas....

by the absence of the corrugated rims which surround each mesh of the C. crispus fruit body. The phylogenetically close species C. chrysomycelinus
Clathrus chrysomycelinus
Clathrus chrysomycelinus is a species of fungus in the stinkhorn family. It is found in South America....

has a yellow receptacle with arms that are structurally simpler, and its gleba is concentrated on specialized "glebifers" located at the lattice intersections. It is known only from Venezuela to southern Brazil. Clathrus columnatus
Clathrus columnatus
Clathrus columnatus, commonly known as the column stinkhorn, is a saprobic species of basidiomycete fungus in the family Phallaceae. It has a widespread distribution, and has been found in Africa, Australasia, and the Americas. It may have been introduced to North America with exotic plants...

has a fruit body with two to five long vertical orange or red spongy columns, joined together at the apex.

Edibility and folklore

Although edibility
Edible mushroom
Edible mushrooms are the fleshy and edible fruiting bodies of several species of fungi. Mushrooms belong to the macrofungi, because their fruiting structures are large enough to be seen with the naked eye. They can appear either below ground or above ground where they may be picked by hand...

 for C. ruber has not been officially documented, its foul smell would dissuade most individuals from consuming it. In general, stinkhorn mushrooms are considered edible when still in the egg stage, and are even considered delicacies in some parts of Europe and Asia, where they are pickled raw and sold in markets as "devil's eggs". However, an 1854 report provides a cautionary tale to those considering consuming the mature fruit body. Dr. F. Peyre Porcher, of Charleston, South Carolina
Charleston, South Carolina
Charleston is the second largest city in the U.S. state of South Carolina. It was made the county seat of Charleston County in 1901 when Charleston County was founded. The city's original name was Charles Towne in 1670, and it moved to its present location from a location on the west bank of the...

, described an account of poisoning caused by the mushroom:
"A young person having eaten a bit of it, after six hours suffered from a painful tension of the lower stomach, and violent convulsions. He lost the use of his speech, and fell into a state of stupor, which lasted for forty-eight hours. After taking an emetic he threw up a fragment of the mushroom, with two worms, and mucus, tinged with blood. Milk, oil, and emollient fomentations, were then employed with success."


British mycologist Donald Dring, in his 1980 monograph
Monograph
A monograph is a work of writing upon a single subject, usually by a single author.It is often a scholarly essay or learned treatise, and may be released in the manner of a book or journal article. It is by definition a single document that forms a complete text in itself...

 on the Clathraceae family, wrote that C. ruber was not regarded highly in southern European folklore. He mentions a case of poisoning following its ingestion, reported by Barla
Jean-Baptiste Barla
Jean-Baptiste Barla was a French botanist.Barla was a man of independent means and dedicated himself to investigate the fungi and orchids of the South of France. "Cette végétation exceptionelle, doit évidemment sa richesse et sa variété à la configuration topographique, toute particulière, du...

 in 1858, and notes that Ciro Pollini reported finding it growing on a human skull in a tomb in a deserted church. According to John Ramsbottom
John Ramsbottom (mycologist)
John Ramsbottom was a British mycologist.He was Keeper of Botany at the British Museum . He served as general secretary and twice as president of the British Mycological Society, and was long editor of its Transactions. He was president of the Linnean Society from 1937 to 1940 and was awarded...

, Gascons
Gascony
Gascony is an area of southwest France that was part of the "Province of Guyenne and Gascony" prior to the French Revolution. The region is vaguely defined and the distinction between Guyenne and Gascony is unclear; sometimes they are considered to overlap, and sometimes Gascony is considered a...

 consider the mushroom a cause of cancer; they will usually bury specimens they find. In other parts of France it has been reputed to produce skin rashes
Eczema
Eczema is a form of dermatitis, or inflammation of the epidermis . In England, an estimated 5.7 million or about one in every nine people have been diagnosed with the disease by a clinician at some point in their lives.The term eczema is broadly applied to a range of persistent skin conditions...

 or cause convulsions.

Ecology, habitat, and distribution

Like most of the species of the Phallales
Phallales
The Phallales are an order of fungi that is more or less synonymous with the gomphoid-phalloid clade. The order contains two families, the Claustulaceae and the Phallaceae, which collectively contain 26 genera and 88 species.-External links:...

 order, Clathrus ruber is saprobic—a decomposer of wood and plant organic matter—and is commonly found fruiting in mulch
Mulch
In agriculture and gardening, is a protective cover placed over the soil to retain moisture, reduce erosion, provide nutrients, and suppress weed growth and seed germination. Mulching in gardens and landscaping mimics the leaf cover that is found on forest floors....

 beds. The fungus grows alone or clustered together near woody debris, in lawns, gardens, and cultivated soil.

Clathrus ruber was originally described by Micheli from Italy. It is considered native to southern and central continental Europe, as well as Macaronesia
Macaronesia
Macaronesia is a modern collective name for several groups of islands in the North Atlantic Ocean near Europe and North Africa belonging to three countries: Portugal, Spain, and Cape Verde...

 (the Azores
Azores
The Archipelago of the Azores is composed of nine volcanic islands situated in the middle of the North Atlantic Ocean, and is located about west from Lisbon and about east from the east coast of North America. The islands, and their economic exclusion zone, form the Autonomous Region of the...

 and the Canary Islands
Canary Islands
The Canary Islands , also known as the Canaries , is a Spanish archipelago located just off the northwest coast of mainland Africa, 100 km west of the border between Morocco and the Western Sahara. The Canaries are a Spanish autonomous community and an outermost region of the European Union...

), western Turkey, North Africa (Algeria), and western Asia (Iran). The fungus is rare in central Europe, and is listed in the Red data book
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

 of Ukraine.

The fungus has probably been introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 elsewhere, often because of the use of imported mulch used in gardening and landscaping
Landscaping
Landscaping refers to any activity that modifies the visible features of an area of land, including:# living elements, such as flora or fauna; or what is commonly referred to as gardening, the art and craft of growing plants with a goal of creating a beautiful environment within the landscape.#...

. It may have extended its range northwards into the British Isles or been introduced in the nineteenth century. It now has a mainly southerly distribution in England and has been recorded from Cornwall
Cornwall
Cornwall is a unitary authority and ceremonial county of England, within the United Kingdom. It is bordered to the north and west by the Celtic Sea, to the south by the English Channel, and to the east by the county of Devon, over the River Tamar. Cornwall has a population of , and covers an area of...

, Devon
Devon
Devon is a large county in southwestern England. The county is sometimes referred to as Devonshire, although the term is rarely used inside the county itself as the county has never been officially "shired", it often indicates a traditional or historical context.The county shares borders with...

, Dorset
Dorset
Dorset , is a county in South West England on the English Channel coast. The county town is Dorchester which is situated in the south. The Hampshire towns of Bournemouth and Christchurch joined the county with the reorganisation of local government in 1974...

, Somerset
Somerset
The ceremonial and non-metropolitan county of Somerset in South West England borders Bristol and Gloucestershire to the north, Wiltshire to the east, Dorset to the south-east, and Devon to the south-west. It is partly bounded to the north and west by the Bristol Channel and the estuary of the...

, the Isle of Wight
Isle of Wight
The Isle of Wight is a county and the largest island of England, located in the English Channel, on average about 2–4 miles off the south coast of the county of Hampshire, separated from the mainland by a strait called the Solent...

, Hampshire
Hampshire
Hampshire is a county on the southern coast of England in the United Kingdom. The county town of Hampshire is Winchester, a historic cathedral city that was once the capital of England. Hampshire is notable for housing the original birthplaces of the Royal Navy, British Army, and Royal Air Force...

, Sussex
Sussex
Sussex , from the Old English Sūþsēaxe , is an historic county in South East England corresponding roughly in area to the ancient Kingdom of Sussex. It is bounded on the north by Surrey, east by Kent, south by the English Channel, and west by Hampshire, and is divided for local government into West...

, Surrey
Surrey
Surrey is a county in the South East of England and is one of the Home Counties. The county borders Greater London, Kent, East Sussex, West Sussex, Hampshire and Berkshire. The historic county town is Guildford. Surrey County Council sits at Kingston upon Thames, although this has been part of...

, and Middlesex
Middlesex
Middlesex is one of the historic counties of England and the second smallest by area. The low-lying county contained the wealthy and politically independent City of London on its southern boundary and was dominated by it from a very early time...

. In Scotland, it has been recorded from Argyll
Argyll
Argyll , archaically Argyle , is a region of western Scotland corresponding with most of the part of ancient Dál Riata that was located on the island of Great Britain, and in a historical context can be used to mean the entire western coast between the Mull of Kintyre and Cape Wrath...

. It is also known from Wales, the Channel Islands
Channel Islands
The Channel Islands are an archipelago of British Crown Dependencies in the English Channel, off the French coast of Normandy. They include two separate bailiwicks: the Bailiwick of Guernsey and the Bailiwick of Jersey...

, and Ireland. The fungus also occurs in the United States (California
California
California is a state located on the West Coast of the United States. It is by far the most populous U.S. state, and the third-largest by land area...

, Florida
Florida
Florida is a state in the southeastern United States, located on the nation's Atlantic and Gulf coasts. It is bordered to the west by the Gulf of Mexico, to the north by Alabama and Georgia and to the east by the Atlantic Ocean. With a population of 18,801,310 as measured by the 2010 census, it...

, Georgia, Virginia
Virginia
The Commonwealth of Virginia , is a U.S. state on the Atlantic Coast of the Southern United States. Virginia is nicknamed the "Old Dominion" and sometimes the "Mother of Presidents" after the eight U.S. presidents born there...

, North Carolina
North Carolina
North Carolina is a state located in the southeastern United States. The state borders South Carolina and Georgia to the south, Tennessee to the west and Virginia to the north. North Carolina contains 100 counties. Its capital is Raleigh, and its largest city is Charlotte...

, and New York
New York
New York is a state in the Northeastern region of the United States. It is the nation's third most populous state. New York is bordered by New Jersey and Pennsylvania to the south, and by Connecticut, Massachusetts and Vermont to the east...

), Canada, Mexico, and Australasia. The species was also reported from South America (Argentina). In China, it has been collected from Guangdong
Guangdong
Guangdong is a province on the South China Sea coast of the People's Republic of China. The province was previously often written with the alternative English name Kwangtung Province...

, Sichuan
Sichuan
' , known formerly in the West by its postal map spellings of Szechwan or Szechuan is a province in Southwest China with its capital in Chengdu...

, Guizhou
Guizhou
' is a province of the People's Republic of China located in the southwestern part of the country. Its provincial capital city is Guiyang.- History :...

, and Tibet
Tibet
Tibet is a plateau region in Asia, north-east of the Himalayas. It is the traditional homeland of the Tibetan people as well as some other ethnic groups such as Monpas, Qiang, and Lhobas, and is now also inhabited by considerable numbers of Han and Hui people...

. Records from Japan are referable to Clathrus kusanoi
Clathrus kusanoi
Clathrus kusanoi is a species of fungus in the stinkhorn family. It is known only from Japan....

; records from the Caribbean are probably of C. crispus.

Biochemistry

Like other stinkhorn fungi, C. ruber bioaccumulates
Bioaccumulation
Bioaccumulation refers to the accumulation of substances, such as pesticides, or other organic chemicals in an organism. Bioaccumulation occurs when an organism absorbs a toxic substance at a rate greater than that at which the substance is lost...

 the element manganese
Manganese
Manganese is a chemical element, designated by the symbol Mn. It has the atomic number 25. It is found as a free element in nature , and in many minerals...

. It has been postulated that this element plays a role in the enzymatic
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

 breakdown of the gleba with simultaneous formation of odorous compounds. Compounds like dimethyl sulfide
Dimethyl sulfide
Dimethyl sulfide or methylthiomethane is an organosulfur compound with the formula 2S. Dimethyl sulfide is a water-insoluble flammable liquid that boils at and has a characteristic disagreeable odor. It is a component of the smell produced from cooking of certain vegetables, notably maize,...

, aldehyde
Aldehyde
An aldehyde is an organic compound containing a formyl group. This functional group, with the structure R-CHO, consists of a carbonyl center bonded to hydrogen and an R group....

s, and amine
Amine
Amines are organic compounds and functional groups that contain a basic nitrogen atom with a lone pair. Amines are derivatives of ammonia, wherein one or more hydrogen atoms have been replaced by a substituent such as an alkyl or aryl group. Important amines include amino acids, biogenic amines,...

s—which contribute to the disagreeable odor of the gleba—are produced by the enzymatic decarboxylation
Decarboxylation
Decarboxylation is a chemical reaction that releases carbon dioxide . Usually, decarboxylation refers to a reaction of carboxylic acids, removing a carbon atom from a carbon chain. The reverse process, which is the first chemical step in photosynthesis, is called carbonation, the addition of CO2 to...

 of keto acid
Keto acid
Keto acids are organic compounds that contain a carboxylic acid group and a ketone group. The alpha-keto acids are especially important in biology as they are involved in the Krebs citric acid cycle and in glycolysis...

s and amino acid
Amino acid
Amino acids are molecules containing an amine group, a carboxylic acid group and a side-chain that varies between different amino acids. The key elements of an amino acid are carbon, hydrogen, oxygen, and nitrogen...

s, but the enzymes will only work in the presence of manganese. A chemical analysis
Elemental analysis
Percent Composition is a process where a sample of some material is analyzed for its elemental and sometimes isotopic composition. Elemental analysis can be qualitative , and it can be quantitative...

 of the elemental
Chemical element
A chemical element is a pure chemical substance consisting of one type of atom distinguished by its atomic number, which is the number of protons in its nucleus. Familiar examples of elements include carbon, oxygen, aluminum, iron, copper, gold, mercury, and lead.As of November 2011, 118 elements...

 composition of the gelatinous outer layer, the embryonic receptacle and the gleba showed the gelatinous layer to be richest in potassium
Potassium
Potassium is the chemical element with the symbol K and atomic number 19. Elemental potassium is a soft silvery-white alkali metal that oxidizes rapidly in air and is very reactive with water, generating sufficient heat to ignite the hydrogen emitted in the reaction.Potassium and sodium are...

, calcium
Calcium
Calcium is the chemical element with the symbol Ca and atomic number 20. It has an atomic mass of 40.078 amu. Calcium is a soft gray alkaline earth metal, and is the fifth-most-abundant element by mass in the Earth's crust...

, manganese, and iron
Iron
Iron is a chemical element with the symbol Fe and atomic number 26. It is a metal in the first transition series. It is the most common element forming the planet Earth as a whole, forming much of Earth's outer and inner core. It is the fourth most common element in the Earth's crust...

 ions. Calcium2+ stabilizes the polysaccharide
Polysaccharide
Polysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...

 gel, protecting the embryonic receptacle from drying out during the growth of the egg. Potassium is required for the gelatinous layer to retain its osmotic pressure
Osmotic pressure
Osmotic pressure is the pressure which needs to be applied to a solution to prevent the inward flow of water across a semipermeable membrane....

 and retain water; high concentrations of the element are needed to support the rapid growth of the receptacle. The high concentration of elements suggests that the gelatinous layer has a "placenta
Placenta
The placenta is an organ that connects the developing fetus to the uterine wall to allow nutrient uptake, waste elimination, and gas exchange via the mother's blood supply. "True" placentas are a defining characteristic of eutherian or "placental" mammals, but are also found in some snakes and...

-like" function—serving as a reservoir from which the receptacle may draw upon as it rapidly expands.

Pigment
Pigment
A pigment is a material that changes the color of reflected or transmitted light as the result of wavelength-selective absorption. This physical process differs from fluorescence, phosphorescence, and other forms of luminescence, in which a material emits light.Many materials selectively absorb...

s responsible for the orange to red colors of the mature fruit bodies have been identified as carotene
Carotene
The term carotene is used for several related unsaturated hydrocarbon substances having the formula C40Hx, which are synthesized by plants but cannot be made by animals. Carotene is an orange photosynthetic pigment important for photosynthesis. Carotenes are all coloured to the human eye...

s, predominantly lycopene
Lycopene
Lycopene is a bright red carotene and carotenoid pigment and phytochemical found in tomatoes and other red fruits and vegetables, such as red carrots, watermelons and papayas...

 and beta-carotene—the same compounds responsible for the red and orange colors of tomatoes and carrots, respectively. Lycopene is also the main pigment in the closely related fungus Clathrus archeri
Clathrus archeri
Clathrus archeri , commonly known as Octopus Stinkhorn, is indigenous to Australia and Tasmania and an introduced species in Europe and North America. The young fungus erupts from a suberumpent egg by forming into four to seven elongated slender arms initially erect and attached at the top...

, while beta-carotene is the predominant pigment in the Phallaceae species Mutinus caninus
Mutinus caninus
Mutinus caninus, commonly known as the dog stinkhorn, is a small thin, phallus-shaped woodland fungus, with a dark tip. It is often found growing in small groups on wood debris, or in leaf litter, during summer and autumn in Europe, Asia, and eastern North America...

, M. ravenelii
Mutinus ravenelii
Mutinus ravenelii is a species of fungus that is often confused with M. elegans and M. caninus. M. ravenelii is a member of the Phallaceae family.-Edibility:...

, and M. elegans
Mutinus elegans
Mutinus elegans, commonly known as the elegant stinkhorn, the dog stinkhorn, the headless stinkhorn, or the devil's dipstick, is a species of fungus in the Phallaceae family. A saprobic species, it is typically found growing on the ground singly or in small groups on woody debris or leaf litter,...

.

External links

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